Ultrastructural observations on sensory cells and the peripheral nervous system in the mantle edge of Helisoma duryi (Mollusca: Pulmonata)
The dorsal surface of the mantle edge produces the overlying periostracum and shell; the ventral epithelium is not involved in shell formation. This regionalization is reflected in the distribution of epithelial receptors. Six ultrastructurally distinct receptor types have been identified on the ventral surface, particularly at the tip of the mantle edge, and one on the dorsal surface. Dorsal receptors occur in a single row along the edge of the mantle edge gland and are also scattered over the dorsal epithelium. Each receptor has a ciliated intraepithelial segment (receptor ending), connected to the subepithelial cell body by a narrow process, and an axon. Variation occurs in the general morphology of the receptor ending, the number of cilia, and the shape of the ciliary tuft. In most types the cell body and axon contain electron-dense granules. Ventral and dorsal receptors differ in the complexity of their subepithelial connections. Ventral receptors send their axons to branches of the pallial nerves. Intrinsic neurons are present in the pallial nerves and in connective tissue and may be responsible for neuromuscular innervation in the mantle edge. Dorsal receptors synapse with intrinsic neurons in a discrete, localised subepithelial plexus. Three types of intrinsic cell have been identified in the plexus, and synapses between them are common. Intrinsic cells form neuromuscular synapses near the plexus and may also innervate epithelial cells in regions responsible for periostracum thickening and calcium deposition.